New Hippopotamid Finds in Eurotas Valley (Laconia, Greece)

Scientific Annals, School of Geology, Aristotle University of Thessaloniki Thessaloniki Special volume 99 57-62 Proceedings of the XIX CBGA Congress, Thessaloniki, Greece 2010

NEW HIPPOPOTAMID FINDS IN EUROTAS VALLEY (LACONIA, GREECE)

Athanassiou A. and Bouzas D.

Hellenic Ministry of Culture, Department of Palaeoanthropology–Speleology, Ardittou 34B, 116 36 Athens, Greece. [email protected],[email protected]

Abstract: A new locality tracked down in the Eurotas Valley (Laconia, Greece) yielded mammalian dental remains of a young individual referred to as Hippopotamus antiquus. The findings are of very large size compared to already known specimens from Greece and W. Europe. The new locality is biochronologically dated at the Early–Middle Pleistocene. Keywords: Hippopotamus Laconia, Peloponnesus, Greece, Pleistocene

1. Introduction area of the Eurotas Valley: Georgalas (1941) de- scribes a proboscidean molar from Skoúra, SE of During a geological survey in Eurotas valley (La- Sparta, which he refers to the Pliocene species conia, Peloponnesus) in October 2006, a new Anancus arvernensis. Sidiropoulou (1972) refers mammal locality was discovered. The site, dubbed an equid maxillary fragment from Vrontamás, pre- ―Myrtiá‖ (MYP), is located in the area among the serving cheek teeth in both sides, to Equus cabal- small town of Vlachiótis and the villages of Myrtiá lus forma primitiva; however, its dental characters, and Peristéri, near the road that leads from Myrtiá as the short protocones and the narrow styles, indi- to the main road of Skála–Vlachiótis (Fig. 1). It is cate that the specimen belongs to a stenonid horse a small remnant of an old terrace of the river Euro- of Late Pliocene or Early Pleistocene age. In a ra- tas, which outcrops above the present floodplain, vine of the same region Symeonidis (1970) men- forming a low hill (highest altitude: 38 m). It con- tions the presence of a (proboscidean?) tusk frag- sists of gravels and loose conglomerates deposited ment. The broader area of Vrontamás has recently in a fluvial environment. Part of the gravels has yielded some new mammal remains (Skourtsos been quarried several years ago. pers. com. and personal field data). Symeonidis & Theodorou (1986a) describe proboscidean skeletal and dental material from Vlachiótis, in the vicinity of Myrtiá, which they refer to Mammuthus meridionalis. Other Laconian localities, outside the Eu- rotas Valley, include the cave of Dyrós, which is particularly rich in Hippopotamus (Giannopoulos 2000; Symeonidis & Giannopoulos 2001), as well as the important anthropological sites of Apídima (Pitsios 1999; Tsoukala 1999) and Lakonís (Har- Fig. 1. Geographic location of the new fossil mammal vati et al. 2003). locality Myrtiá (MYP). Graphical scale in km. Contour interval: 100 m. 2. Systematics Order: Artiodactyla OWEN, 1848 The new locality yielded two Hippopotamus mo- Family: Hippopotamidae GRAY, 1821 lars that plausibly belong to the same individual, as Genus: Hippopotamus LINNAEUS, 1758 they were found in association and exhibit similar Hippopotamus antiquus DESMAREST, 1822 degree of wear. Material: MYP-1: upper left second molar (M2); Fossil mammal localities of Pliocene to Early MYP-2: upper left third molar (M3) (Fig. 2). The Pleistocene age are already known in the broader material is currently stored in the Museum of Pa-

57 laeontology and Geology of the National and Ka- the lingual and labial openings of the transverse podistrian University of Athens. valleys (the lingual one being more robust). In M3 the cingulum is preserved only mesially and distally. The distal one is weaker than the corresponding structure of M2. The dimensions of M2 are (63.0) × (58.5) × 60.0 (mesiodistal diameter × labiolingual diameter × height, in mm), while those of M3 are (61.0) × (54.0) × 56.8 respectively (the dimensions in parentheses indicate inaccurate measure- ments, due to incomplete preservation). The degree of dental wear can provide information on the ontogenetic age of the Myrtiá individual. According to Laws (1968), who described the age groups of extant African hippos based on the erup- tion sequence of lower teeth, the first signs of wear on lower second molar appear at the age groups VII-VIII. The same author, based though on li- mited observations, states that the same criteria approximately apply on the upper teeth too. Thus the individual of Myrtiá can be aged into these Fig. 2. Hippotamus antiquus from Myrtiá. Left upper second molar (MYP-1): a, occlusal view; b, labial view. groups, which correspond to an ontogenetic age of Left upper third molar (MYP-2): c, occlusal view; d, la- 11±2 to 15±2 years in the extant populations bial view. Mesial side is on the left. Graphical scale in (Laws, 1968). cm. 3. Discussion

Description: The two molars were found in ana- An exclusively African genus during the Holocene, tomical association; however no maxillary bone Hippopotamus was common in Europe during not even the roots are preserved. Both teeth are in most of the Pleistocene (especially in the western pristine condition: the enamel is wrinkled and only part), where it is present in hundreds of palaeonto- the mesial cusps (protocone and paracone) of M2 logical localities (Faure, 1981). The genus oc- show some signs of polishing due to very early curred in South, Central and Western Europe dur- enamel wear. The M3 must not have been erupted ing the Early and early Middle Pleistocene, but at the time of death. In occlusal view the teeth ex- subsequently retreated to the South (Iberian, Italian hibit the typical trefoil-shaped crown morphology, and Balkan peninsulas) till the end of the Middle which is characteristic for the Hippopotamidae. Pleistocene. In the Late Pleistocene the hippos The M2 has trapezoid shape, the mesial cusps be- were again widespread in Western Europe, reach- ing considerably wider than the distal ones. In M3 ing as far north as N. England. In general their the cusp pairs are placed rather oblique in relation biogeographic range extended north during the in- to the sagittal plane. All cusps are conical and con- terglacials and contracted to southern refugia dur- verge towards their apices. In labial or lingual view ing the glacials. The European population dimi- the mesial cusps appear almost straight, or bend nished and eventually became extinct during the slightly backwards, while the distal cusps are last glacial. clearly bent toward the centre of the crown. The mesial and distal cusp pairs are separated by very The earlier forms (Early to Middle Pleistocene), deep transverse valleys that open lingually and la- usually referred to as Hippopotamus antiquus, bially. The styles situated mesially and distally of were very large. Later forms (generally after the each cusp are very strong in M2, particularly the mid-Pleistocene) were smaller, comparable in size hypostyles. In M3 they are considerably weaker; to the recent African species H. amphibius and the distal metastyle is practically lacking, being usually referred to as such. A number of other spe- only visible near the base of the crown. Both mo- cies were also proposed, as H. incognitus FAURE, lars have a cingulum. In M2 it is very strong in all 1984 and H. tiberinus MAZZA, 1991, based on ana- sides of the tooth: mesially and distally it reaches tomical peculiarities of the skull or postcranial ske- the middle of the crown, while it forms a cuspid in leton, though they are not generally accepted with-

58 out reservation (Mazza 1995; Petronio 1995). Oth- er authors consider all European Pleistocene hippopotamuses as belonging to the gradually evolv- ing extant species H. amphibius, in which they q recognize two subspecies (H. amphibius antiquus e w r and H. amphibius amphibius) (e.g. Kahlke, 1990; ty Kahlke, 2001). Since the taxonomy of the genus at u the species/subspecies level is still debated, the two species (antiquus–amphibius) taxonomic i scheme is provisionally accepted in the present 1@ study, following Petronio (1995). o 1# 1)1! 1$1^ Non-endemic Hippopotamus is already known 1% 2) from several sites in Greece (Fig. 3), but it is most- 1( 1& ly represented by few fragmentary finds in each 1* site. The richest sites are located in the Megalópo- lis Basin —sites Megalópolis (Melentis, 1966a) and Kyparíssia (personal data)—, as well as in

Dyrós Cave (Petrochilos, 1958; Giannopoulos, Fig. 3. Distribution of non-endemic Hippopotamus- 2000; Symeonidis and Giannopoulos, 2001). The bearing localities in Greece: 1, Ravin Voulgárakis, presence of Hippopotamus sp. is also mentioned 2, Haliákmon Valley (unknown exact location), 3, Per- by Deprat (1904) at Panagia Heria, Euboea, to- díkkas, 4, Libákos, 5, Kapetánios, 6, Q-Profil, 7, Piniós gether with a Late Miocene assemblage, which Valley, 8, Toíchos, 9, Manzavináta, 10, Elis, 11, Ag. makes this determination rather doubtful. In a later Dimítrios, 12, Káto Salmeníko, 13, Limnón Cave, publication on the Miocene faunas of the island 14, Kyparíssia, 15, Megalópolis, 16, Marathoúsa, (Mitzopoulos, 1947) the genus is not included in 17, Apídima Caves, 18, Dirós Cave, 19, Myrtiá, the faunal lists. Endemic Hippopotamus (H. 20, Kos. Data according to Desio (1931), Psarianos (1954), Thenius (1955), Milójčić et al. (1965), Melentis creutzburgi BOEKSCHOTEN & SONDAAR, 1966) is (1966a, 1966b, 1969), Sickenberg (1976), Stratigopou- currently known from twenty localities in Crete los (1986), Symeonidis and Theodorou 1986b) Steen- (Lax, 1996; Iliopoulos et al., 2010), the most im- sma (1988), Koufos et al. (1989), Tsoukala (1999), portant of which is in the mountain basin of Ka- Symeonidis and Giannopoulos (2001), Reimann and tharó (Eastern Crete) (Boekschoten and Sondaar, Strauch (2008) and personal observations. 1966; Dermitzakis et al., 2005). as well as those from Dyrós, are much smaller than The Hippopotamus molars have only minimally Myrtiá, which is also consistent with their specific changed during the genus’ evolution and, conse- determination by the above authors as H. amphi- quently, their morphology is not taxonomically di- bius. The two M2s from Dyrós (specimens N.84 agnostic. Nevertheless, the trapezoid outline of M2 and N.86; Museum of Palaeontology and Geology, in occlusal view has been considered as a specific University of Athens) are, however, close to the character of H. antiquus (Thenius, 1955). In a me- maximum values of the species’ range, as given by trical comparison to other dental samples of Euro- Faure (1985). The M2 from Elis is similar in size pean Hippopotamus (Fig 4), the molars from Myr- to MYP-1, while the M2 from Káto Salmeníko is tiá plot near the maxima of H. antiquus ranges ac- slightly smaller. The locality of Ravin Voulgarákis cording to Faure (1985) and appear to be among has yielded an almost complete juvenile skull that the largest specimens of the same species accord- preserves the M2s. With a mesiodistal dimension ing to Mazza (1995). Hippopotamus upper molars of about 55 mm (Kostopoulos, 1996, Table 146) from Greece, which can be metrically compared to the M2s of this skull are smaller than the ones the Myrtiá material, are known from the following from Myrtiá. localities: Manzavináta (Psarianos, 1954), Elis (Thenius, 1955), Haliákmon Valley (Melentis The available morphological and —especially— 1966b), Káto Salmeníko (Symeonidis and Theodo- metrical characters of the molars from Myrtiá sup- rou, 1986b), Ravin Voulgarákis (Kostopoulos, port the attribution of the finds to the species Hip- 1996) and Dyrós Cave (Giannopoulos, 2000). The popotamus antiquus. molars from Manzavináta and Haliákmon Valley,

70 with mollusk faunas, and this age has been recently M2

65 corroborated by Reimann and Strauch (2008), who studied a partially preserved skull from the same 60 area. This places the finds from Elis among the

55 earliest Hippopotamus of Europe. The other hippo- yielding localities of Greece are of later or not well 50 definable age. Steensma (1988) dates the localities

45 of Libákos and Kapetánios to the biozone MNQ19

labiolingualdiameter (Early Pleistocene). Slightly younger, Ravin Voul- 40 garákis is dated by Koufos (2001) in MNQ20. The

35 findings of the Megalópolis Basin (sites Me- 30 35 40 45 50 55 60 65 70 galópolis, Marathousa and Kyparíssia) are of Mid- m e s i o d i s t a l d i a m e t e r dle Pleistocene age, as the lacustrine sediments of 65 M3 the basin are dated from ~900 ka to ~350 ka (van Vugt et al., 2000). Of the three faunas, Marathousa 60 is probably the older, about at the Early/Middle

55 Pleistocene boundary, while Kyparíssia is possibly closer to the top of the stratigraphic sequence. 50 The fauna of Piniós Valley is dated by Milójčić et

45 al. (1965) to the last interglacial or the beginning

labiolingualdiameter of the last glacial period. However, radiometric 40 dates cited by Runnels & van Andel (1999) place the fossiliferous sediments well into the late gla- 35 40 45 50 55 60 65 70 cial. If these late dates (~ 45–30 ka) are corrobo- m e s i o d i s t a l d i a m e t e r rated, then the taxonomic attribution of the finds, Myrtiá Elis Haliákmon Manzavináta Dyrós Káto Salmeníko referred by Milójčić et al. (1965) to as H. am- H. antiquus H. amphibius Valdarno phibius cf. antiquus, might need revision. New ma- Fig. 4. Scatter plots comparing the mesiodistal (DAP) to terial collected recently at the locality may yield labiolingual (DT) dimensions in mm of Hippopotamus additional data on this problem (Athanassiou in M2 and M3 samples. The blue and green squares repre- preparation). A late date would also raise the upper sent the range given by (Faure 1985) for H. amphibius biochronological limit of the genus in Europe, and H. antiquus respectively. The blue and green points which is currently placed at Oxygen Isotope Stage correspond to H. amphibius and H. antiquus measure- 5 or 4 (Petronio, 1995; Palombo and Mussi, 2001). ments respectively, according to Mazza (1995). Meas- A late radiometric date of at least 32 ka is also urements of Valdarno, Manzavináta, Elis, Haliákmon Valley and Káto Salmeníko according to Leonardi cited by Symeonidis and Giannopoulos (2001) for (1948), Psarianos (1954), Thenius (1955), Melentis the H. amphibius-bearing fauna of Dyrós Cave. (1966b) and Symeonidis and Theodorou (1986b). The The specimens from Manzavináta and Haliákmon plots depict clearly the great metrical overlapping of Valley are attributed by Melentis (1966b) and Psa- samples referred to as H. amphibius and H. antiquus, rianos (1953) to H. amphibius, on the base of their which leads to confusion regarding the taxonomy of the European representatives of the genus. rather smaller dimensions, so a Middle–Late Pleis- tocene age is more probable for these localities. A 4. Biochronology–Palaeoecology stratigraphic correlation of several European Hip- popotamus-bearing localities is given in the chart Azzaroli et al. (1982) place the first appearance of of figure 5. Hippopotamus in Europe at the Early Pleistocene (Tasso Faunal Unit, at about 1.8–1.7 Ma), as it is The stratigraphic range of H. antiquus is usually known during this time interval from localities in given as Early–Middle Pleistocene (Faure, 1981; Italy and other Mediterranean regions. However, Mazza, 1995; Petronio, 1995) and this is a plausi- scanty specimens from sites of rather earlier, Late ble age for the locality of Myrtiá. Given the very Pliocene age make probable an even earlier disper- large dimensions of the studied specimens and the sal from Africa to Europe (Kahlke, 1987). Thenius suggested Late Pliocene age of the Elis finds, a (1955) dates his H. antiquus findings from Elis to similar age could also be supposed as possible for the Astian (Late Pliocene), based on correlations Myrtiá. However, since Hippopotamus remains

60 have not yet been found in any well-dated Pliocene to fossils already known from several European lo- mammal assemblage in Greece, we reserve a Late calities. The presence of Hippopotamus in Myrtiá Pliocene date, until the early presence of the genus suggests a fluvial environment. The new locality is in Greece is supported with more biostratigraphic biochronologically dated at the Early–Middle data from more sites. The presence of Mammuthus Pleistocene. meridionalis in the nearby locality of Vlachiótis (Symeonidis and Theodorou 1986a) also indicates Acknowledgements a wide chronological range of Late Pliocene – Ear- The authors wish to thank Paris Pavlakis and an ly Pleistocene. A direct stratigraphic correlation anonymous reviewer, whose suggestions improved between the two localities is, though, not possible. the paper.

Geomag. Mammal References Ma scale Epochs biochronology L o c a l i t i e s

HOLOCENE MNQ 26 Dyrós Cave late Piniós Valley MNQ 25 Azzaroli A., De Giuli C., Ficcarelli G. and Torre D., Barrington 1982. Table of the stratigraphic distribution of terres- MNQ 24

AURELIAN trial mammalian faunas in Italy from the Pliocene to MNQ 23 Kyparíssia (?) MNQ 22 0.5 H . a m p h i b i u s Mosbach the early middle Pleistocene. Geografia Fisica e Di- m i d d l e namica Quaternaria, 5, 55-58. BRUNHES MNQ 21 Mauer Forest Bed Megalópolis Isernia Akhalkalaki Boekschoten G.J. and Sondaar P.Y., 1966. The Pleisto- Durfort GALERIAN cene of the Katharo Basin (Crete) and its Hippopota- mus. Bijdragen tot de Dierkunde, 36, 17-44. 1.0 MNQ 20 Untermaßfeld Saint-Prest Colle Curti Deprat M.J., 1904. Étude géologique et pétrographique

Ravin Voulgarákis de l’île d’Eubée. Besançon, Paris, 231p. Dermitzakis M., Pavlakis P., Drinia H., 2005. The Pleis- e a r l y tocene hippopotamuses of the Katharó Plateau, La-

1.5 l a t e

H i p p o p o t a m u s a n t i q u u s Upper Valdarno Libákos sithi, Crete. Preliminary report. Proceedings of the PLEISTOCENEMNQ 19 Poggio ai Venti Kapetánios (?) 2nd Pancretian Speleological Symposium, Herákleion, 53-59. MATUYAMA Olduvai Olduvai Desio A., 1931. Le isole Italiane dell’Egeo. Memorie Elis 2.0 MNQ 18 Descrittive della Carta Geologica d’Italia, 24, 1-534. VILLAFRANCHIAN middle Faure M., 1981. Répartition des Hippopotamidae (Mam- Fig. 5. Stratigraphic chart of selected European Hippo- malia, Artiodactyla) en Europe occidentale. Implica- potamus-bearing localities, including well-dated Greek tions stratigraphiques et paléoécologiques. Geobios, 14, 191-200. sites. Chronostratigraphic and magnetostratigraphic Faure M., 1985. Les Hippopotames quaternaires non- subdivisions according to Gibbard and van Kolfschoten insulaires d’Europe occidentale. Nouvelles Archives (2005). Mammal biozonation according to Guérin du Muséum d’Histoire Naturelle de Lyon, 23, 13-79. (1990) Faunal data references are given in text. Georgalas G.C., 1941. Über das Vorkommen von Anancus (Mastodon) arvernensis CROIZ. und JOB. in Hippos are adapted to a semi-aquatic way of life der Umgebung von Skoura (SÖ von Sparta). Praktiká and depend on the presence of fresh water bodies Akademías Athinón, 16, 94-100. throughout the year. Their presence has also been Giannopoulos V.I., 2000. Contribution to the study of re- interpreted as an indication of warm climate. This cent and old environments of the most important Greek caves. PhD thesis, National and Kapodistrian is, though, not true, at least for southern Europe, as University of Athens, 437p. (in Greek). already pointed out by Faure (1981) and as indi- Gibbard P. and van Kolfschoten T., 2005. The Pleisto- cated by recently available radiometric dates for cene and Holocene Epochs. In: A Geologic Time Piniós Valley and Dyrós Cave, which place these Scale 2004, Gradstein F.M., Ogg J.G. and Smith A.G. faunas into the last glacial. The presence of H. an- (eds), Cambridge University Press, New York, 441- 452. tiquus in Myrtiá, in combination to the coarse- Guérin C., 1990. Biozones or Mammal Units? Methods grained lithology of the site, indicates a fluvial en- and limits in biochronology. In: European Neogene vironment. mammal chronology, Lindsay E.H., Fahlbusch V. and Mein P. (eds), Plenum, New York, 119-130. 5. Conclusion Harvati K., Panagopoulou E. and Karkanas P., 2003. First Neanderthal remains from Greece: the evidence from The dental findings (upper M2 and M3) from the Lakonis. Journal of Human Evolution, 45, 465-473. new locality Myrtiá are referred to a young indi- Iliopoulos G., Eikamp H. and Fassoulas C., 2010. A new vidual of Hippopotamus antiquus. The specific de- late Pleistocene mammal locality from Western Crete. termination is mainly based on the specimens’ very Proceedings of the 12th International Congress of the large size, compared to recent material, as well as Geological Society of Greece, Patras, Bulletin of the Geological Society of Greece, 42.

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